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Modern Human Origins Modern Human CHAPTER 29 Origins Mark Collard and Mana Dembo INTRODUCTION The origin of modern humans has long been one of the most dynamic topics in paleoanthropology. In the last few years, however, there has been a dramatic increase in the amount of data that are relevant to the issue. This is partly due to discoveries made in the course of fieldwork (e.g., Duarte et al., 1999 ; White et al., 2003 ). But mainly it is a consequence of the development of a range of sophisticated laboratory methods that have allowed us to not only date and compare fossils much more pre- cisely and accurately than was possible before, but also access completely new types of data, including – most remarkably – gene sequences from the fossilized remains of individuals who died over 30,000 years ago (e.g., Krings et al., 1997; Spoor et al., 2003 ; Harvati, 2009 ; Richards and Trinkaus, 2009 ; Reich et al., 2010 ). In this chapter, we provide an overview of the current state of research on modern human origins. Conventionally, the evidence pertaining to modern human origins is evaluated in relation to two models – the African replacement model (also known as the Afro-European sapiens model, the out of Africa model, the Eve theory, the recent African origin model, or the replacement model) and the multiregional evolution model (also known as the multiregional continuity model) (e.g., Wolpoff and Caspari, 1997 ; Wood and Baker, 2011 ). However, it has been argued that treating all the mod- els that have been put forward to explain modern human origins as variants of either the African replacement model or the multiregional evolution model is unhelpful, and that additional models should be recognized (Aiello, 1993 ; Stringer, 2001 ). We find this argument convincing, and therefore assess the evidence against several models. A Companion to Paleoanthropology, First Edition. Edited by David R. Begun. © 2013 Blackwell Publishing Ltd. Published 2013 by Blackwell Publishing Ltd. 0001694005.INDD 557 11/9/2012 11:24:21 AM 558 MARK COLLARD AND MANA DEMBO The remainder of this chapter is divided into two main sections. In the first, we outline the four models we think need to be taken into account when discussing modern human origins. In our view, the modern human origins debate has been ham- pered by ambiguities in the way some of the models in question have been characterized. We discuss these ambiguities and then suggest revisions to the models that reduce the uncertainty. In the other section, we review how well the available anatomical, genetic, and linguistic evidence support the predictions of the models. Anatomical, genetic, and linguistic research on modern human origins has tended to focus on three issues: the structure of living human variation, the timing of the appear- ance of modern humans, and evidence for the occurrence of gene flow between mod- ern humans and nonmodern hominins. We discuss each of these in turn. MODELS OF MODERN HUMAN ORIGINS Two papers are particularly helpful for identifying the models that need to be taken into account when discussing modern human origins: Aiello ( 1993 ) and Stringer ( 2001 ). Aiello ( 1993 ) argued that four models could be identified in the literature dealing with modern human origins: the African replacement model, the (African) hybridization and replacement model, the assimilation model, and the multiregional evolution model. These models, she suggested, differ in relation to three factors: geography, timescale, and process. Aiello ( 1993 : 73–74) summarized the models as follows: 1. The African replacement model argues that modern humans first arose in Africa about 100,000 years ago and spread from there throughout the world… Indigenous premodern populations in other areas of the world were replaced by the migrating populations with little, if any, hybridization between the groups. 2. The (African) hybridization and replacement model is similar to the above, but allows for a greater or lesser extent of hybridization among the migrating populations and the indigenous premodern populations. 3. The assimilation model also accepts an African origin for modern humans. However, it differs from the previous models in denying replacement, or population migration, as a major factor in the appearance of modern humans. Rather, this model emphasizes the importance of gene flow, admixture, changing selection pressures, and resulting directional morphological change. In other words, it accepts the fact that, at least in some areas of Eurasia, local evolution (or continuity) could play an important role in the appearance of modern humans. 4. The multiregional evolution model differs from the previous three in denying a recent African origin for modern humans. It emphasizes the role of both genetic continuity over time and gene flow between contemporaneous populations in arguing that modern humans arose not only in Africa but also in Europe and Asia from their Middle Pleistocene forebears. Stringer ( 2001 ) revisited Aiello ’ s ( 1993 ) scheme in light of work published in the 1990s that he believed had caused confusion about the differences among the models. He focused particularly on work that had been claimed to support the multiregional evolution model. In some of the studies in question, he argued, the name “multi- regional evolution model” had been applied to models that should be considered variants of the assimilation model, while in others the name “multiregional evolution 0001694005.INDD 558 11/9/2012 11:24:22 AM MODERN HUMAN ORIGINS 559 model” had been applied to a model that does not appear in Aiello ’ s ( 1993 ) scheme. The latter model, Stringer ( 2001 ) suggested, shares features with the assimilation model and the multiregional evolution model, but is different from both. It is similar to the assimilation model in that it holds that African populations made the largest contribution to the modern human gene pool due to their numerical dominance. Where it differs from the assimilation model and overlaps with the multiregional evolution model is in the timescale involved. Whereas the assimilation model focuses on the Late Pleistocene, the new model contends that the genetic influence of African populations extends throughout the Pleistocene, which is also the timescale the multiregional evolution model operates on. Stringer ( 2001 ) went on to suggest that the new model should be dubbed “multiregional evolution 2” and that the original multiregional evolution model should be renamed “multiregional evolution 1.” Generally, we accept the Aiello ( 1993 ) / Stringer ( 2001 ) scheme. However, we think it needs minor revision. To begin with, the timescale on which the African replacement model and the (African) hybridization and replacement model operate needs to be adjusted. Since the publication of Stringer ( 2001 ), the first appearance date of modern humans in Africa has been pushed back on the basis of new finds and re-dating of previously discovered material to between 160 and 195 ka before the present 1 (White et al., 2003 ; McDougall et al., 2005 ). As such, both models now argue that modern humans first arose in Africa about 200,000 years ago. The second revision we wish to propose concerns the model Stringer ( 2001 ) called “multiregional 2.” It seems to us that this model is better viewed as a variant of the assimilation model than as a version of the multiregional evolution model, and should be renamed accordingly. It is clear from the early work of proponents of the multire- gional evolution model not only that they viewed the genetic contributions of nonmodern hominins 2 from Eurasia and Australasia to modern human populations in those areas to be more important than the genetic contributions of African popula- tions, but also that this is a core component of the model. The following quotes from Thorne and Wolpoff ( 1992 : 78–79) illustrate this. Writing about China, Thorne and Wolpoff make the following claim: “Our examinations of the Chinese specimens found no anatomic evidence that typically African features ever replaced those of the ancient Chinese in these regions. Instead there is a smooth transformation of the ancient population into the living peoples of east Asia.” They make a similar point about the fossil record of Australasia: “The hominid fossils from Australasia (Indonesia, New Guinea and Australia) show a continuous anatomic sequence that is uninterrupted by African migrants at any time.” Given that a dominant contribution from Eurasian and Australasian nonmodern hominins to modern human populations in those regions is central to the multiregional evolution model, it is difficult to see how a model that posits a dominant contribution of African populations to modern human populations in Eurasia and Australasia can be considered to be a variant of the multiregional evolu- tion model. Doing so, in our opinion, effectively strips the multiregional evolution model of one of its key distinguishing features. This problem does not arise if the model that Stringer refers to as multiregional 2 is treated a variant of the assimila- tion model, because the latter assumes that African populations contribute most to the modern human gene pool. Thus, we prefer to call the model “assimilation 2.” Our third proposed revision relates to the distinction between the African replace- ment model and the (African) hybridization and replacement model. In the way these 0001694005.INDD 559 11/9/2012 11:24:22 AM 560 MARK COLLARD AND MANA DEMBO models were described by Aiello ( 1993 ), there is potential
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